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Nomenclature and Taxonomic Status of the Lizards Listed by Philippi (1860) (Squamata: Liolaemidae, Teiidae and Tropiduridae)

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Nomenclature and Taxonomic Status of the Lizards Listed by Philippi (1860) (Squamata: Liolaemidae, Teiidae and Tropiduridae) Trabajo Cuad. herpetol. 35 (Supl. 1): 175-191 (2021) Nomenclature and taxonomic status of the lizards listed by Philippi (1860) (Squamata: Liolaemidae, Teiidae and Tropiduridae) Jaime Troncoso-Palacios1, Yery Marambio-Alfaro2,3 1 Programa de Fisiología y Biofísica, Facultad de Medicina, Universidad de Chile, Independencia 1027, San- tiago, Chile. 2 Laboratorio de Sedimentología y Paleoambientes. Instituto de Ciencias Naturales Alexander von Humboldt, Facultad de Ciencias del Mar y Recursos Biológicos, Universidad de Antofagasta. Avda. Universidad de An- tofagasta 02800, Chile. 3 Parménides Limitada, Avda. Batallones de Atacama 112, Caldera, Atacama, Chile. Recibido: 29 Diciembre 2019 ABSTRACT Revisado: 23 Marzo 2020 In 1860 Rudolph Amandus [Rodolfo Amando] Philippi published the book “Reise durch die Aceptado: 05 Mayo 2020 Wueste Atacama auf Befehl der chilenischen Regierung im Sommer 1853–54”, a wonderful work Editor Asociado: A. S. Quinteros of natural history, which was for a long time the most comprehensive source of geographic, cultural, botanic and zoological information on the Atacama Desert of Chile (currently Atacama and Antofagasta regions). In this book, Philippi listed several species of lizards, and described doi: 10.31017/CdH.2020.(2020-060) five new species. However, the taxonomic identities of several of these species still remain to be clarified. Here we provide comments on these species and show that not all of Philippi`s (1860) type specimens were collected in Atacama Desert as has long been thought. We desig- nate lectotypes for Proctotretus pallidus and P. melanopleurus; and confirm that P. modestus is a junior synonym of L. bellii and that P. pallidus is a junior synonym of L. nigromaculatus. In the case of P. marmoratus, since it is a secondary homonym of L. marmoratus Gravenhorst, 1838, we consider it an invalid name. Key Words: Aporomera, Atacama, Desert, Helocephalus, Liolaemus, Proctotretus. RESUMEN En 1860 Rodolfo Amando [Rudolph Amandus] Philippi publicó el libro (sic) “Viage al Desierto de Atacama hecho de orden del Gobierno de Chile en el verano 1853-54”, un maravilloso tra- bajo de historia natural, el cual fue por mucho tiempo la fuente más completa de información geográfica, cultural, botánica y zoológica del desierto de Atacama de Chile (actualmente las regiones de Atacama y Antofagasta). En este libro, Philippi listo varias especies de lagartos, y describe cinco nuevas especies. Sin embargo, las identidades taxonómicas de varias de estas especies permanecen sin clarificar. Aquí nosotros proveemos comentarios sobre estas especies y demostramos que no todos los especímenes tipo de Philippi (1860) fueron colectados en el desierto de Atacama como se ha pensado por largo tiempo. Nosotros designamos lectotipos para Proctotretus pallidus y P. melanopleurus; y confirmamos que P. modestus es un sinónimo menor de L. bellii y que P. pallidus es un sinónimo menor de L. nigromaculatus. En el caso de P. marmoratus, dado que es un homónimo secundario de L. marmoratus Gravenhorst, 1838, lo consideramos un nombre inválido. Palabras claves: Aporomera, Atacama, Desert, Helocephalus, Liolaemus, Proctotretus. Introduction The German naturalist and explorer Rudolph [Ru- Atacama auf Befehl der chilenischen Regierung im dolfo] Amando Philippi (1808–1904), is well known Sommer 1853–54”, describing his journey and field for his major early contributions to the natural his- survey through the Atacama Desert carried out on tory of Chile. An overview of his life and work was behalf of the Chilean government between Novem- summarized by Kabat and Coan (2017). In 1860 ber 1853 and February 1855. The book provides data Philippi published the book “Reise durch die Wueste on various subjects as geography, culture, botany and Author for correspondence: [email protected] 175 Troncoso-Palacios & Marambio-Alfaro - Lizards listed by Philippi (1860) zoology with a detailed account of the trajectory. group”), which include the nigromaculatus, platei, Philippi`s journey in the Atacama Desert was wide- and other groups; and Eulaemus (or “Argentinean ranging and led from the coast to Andean highlands, group”), which include the darwinii, montanus, and covering the current Atacama and Antofagasta other groups (see Lobo et al., 2010). Also, Philippi regions of northern Chile (Fig. 1). In the Zoology (1860) included species currently assigned to the section of the book (Chapter 8, pp. 156–190), one genus Callopistes, which includes only two species of amphibian and eight lizard species are discussed (pp. larger teiid lizards (Harvey et al., 2012), and from the 165–169), including the description of a new genus to genus Microlophus, which includes approximately and five new species providing a drawing of three of 20 species (Benavides et al., 2007) with several them (reproduced here in Fig. 2). Most of these liz- taxonomic problems among the Chilean species (Troncoso-Palacios, 2018). Almost all species listed by Philippi (1860) are controversial in regard to their taxonomic status or their occurrence in the Atacama Desert (Ortiz and Núñez, 1986). Hereafter, we review the lizards listed by Philippi (1860) on the basis of extant type specimens and comparative material from several Chilean collections and provide com- ments on their nomenclatural and taxonomic status. Materials and methods We examined three type specimens collected by Philippi and an additional seven type specimens of his collection were examined through photographs: Helocephalus nigriceps ZSM 38/1930 lectotype, Proctotretus modestus ZMB 5350, 70546–47, P. mel a - nopleurus FMNH 9969 syntype, and the specimens NMW 18914:1,2 considered by some authors as syntypes of Proctotretus pallidus (Tiedemann and Häupl, 1980; Gemel et al., 2019). For the specimens NMW 18914:1,2 we estimated our measurements based on the rule included in each photograph (in millimeters). We also reviewed photographs of six non type specimens collected by Philippi. For com- parisons, we examined 227 specimens not collected by Philippi but belonging to species that have been Figure 1. Sketch of the route followed by Philippi during his recorded in the area visited by him or in the sur- travel through the Atacama Desert. He started in Coquimbo (1, in November 1853) and traveled by ship to Caldera (3, in May roundings (Appendix I). Scales were observed using 1854), without visit Huasco (2), although some type specimens binocular lenses 0.8–5x. The characters for scalation has been claim as collected in this last locality (see comments were taken according to Pincheira-Donoso and in regards to Proctotretus pallidus). Then, he went to Copiapó (4, in May), returned to Caldera (3, in May). Later he went to Núñez (2005) and Troncoso-Palacios et al. (2015). Chañaral (5, in December), Taltal (6, in December) and Paposo (7, in December), sailed up to Mejillones (8, in December), Results and Conclusion returned to Taltal (6, in January 1855) and traveled to Tilopozo (9, in January) and San Pedro (10, in January). He returned, passing again through Tilopozo (9, in February), Pajonal (11, Proctotretus marmoratus (Philippi 1860, p. 165) in February), Finca de Chañaral (12, in February), and Puquios The genusProctotretus is currently considered a ju- (13, in February) before finally reaching Copiapó (4, in Febru- nior synonym of Stenocercus (Frost, 1992), a genus ary) at the end of his journey. that do not occur in Chile. However, currently all ards currently belong to the genus Liolaemus, which Chilean species formerly listed in the genus Procto- has been split into two subgenera, each split into tretus are placed in the genus Liolaemus (Etheridge, several groups: Liolaemus (sensu stricto or “Chilean 1995; Abdala and Quinteros, 2014), among them, 176 Cuad. herpetol. 35 (Supl. 1): 175-191 (2021) Figure 2. Lizards illustrated in Philippi (1860), here modified from two different copies of the book. Lateral aspect (right), cephalic scalation (center) and dorsal scalation (left). Above=Helocephalus nigriceps. Middle= Proctotretus bisignatus, listed as P. nigromaculatus in the zoological section of Philippi (1860). Below= P. pallidus. P. marmoratus was considered as belonging to Philippi (1860) did not provide additional features Liolaemus by Boulenger (1885, p. 140) who suggest (e.g. color pattern or localities), no conclusion on that it is a questionable junior synonym of L. nitidus the species identity can be reached. (Wiegmann, 1834), an opinion shared by Ortiz and Núñez (1986) and Pincheira-Donoso and Núñez Proctotretus nigromaculatus (Wiegmann, 1834) (2005). Moreover, Philippi (1860) pointed out that (Philippi 1860, p. 166, tab. VI, Fig. 2; reproduced in P. marmoratus “the ventral part from the chin to here in Fig. 2) the anus has rounded and smooth scales”, the smooth Philippi (1860) starts his brief characterization of scales on the throat being a diagnostic trait of Lio- this species (currently Liolaemus nigromaculatus) laemus (Etheridge, 1995). Thus, the correct name attributing the species authority to Wiegmann, but combination is Liolaemus marmoratus (Philippi, then he added the following sentence in regards 1860), however, it is a secondary homonym with L. to the species illustration: “S(ection) Zool(ogy) marmoratus Gravenhorst, 1838 (ICZN, 1999, Art. tab(ularum) nomine Proct(otretus). bisignatus”, 53.3) and then Philippi`s marmoratus is an invalid which introduced a new name (P. bisignatus) as ju- (but available) name. We do not recommend the nior synonym of P. nigromaculatus. The reasons why
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